415
Aspöck 2012) that are parasitic on vertebrates, mostly livestock (Cupp et al. 1998;
Salem et al. 2012), and represent probably a single origination of hematophagy
(Schremmer 1961). Feeding in this tribe is accomplished by sharp, prestomal teeth
on the labellum, resulting in surface abrasion of the integument and sponging of
blood and other dermal fluids that pool in the lesions. The Tribe Stomoxyini lacks a
fossil record, but because the clade is a member of the post-Cretaceous Calyptratae
(Grimaldi and Engel 2005a), the lineage likely originated during the Paleogene and
acquired parasitic interactions with large mammals at that time.
The Hippoboscoidea, termed Pupipara in the older literature, consists of the four
constituent lineages of Glossinidae (tsetse flies), Hippoboscidae (louse flies, keds),
Nycteribiidae (bat flies) and Streblidae (bat flies), although the last two lineages are
not monophyletic (Griffiths 1972; Kutty et al. 2001). Hippoboscoid flies are united
by several, common and distinctive reproductive features. The mature female lays a
single mature larva, the puparium, after which the larva immediately pupariates,
emerging from the hardened, last-instar larval skin that encloses the pupa (Ferrar
1987). The emergent, hematophagous adult then locates and parasitizes a suitable
bird or mammalian host. This unique larval development is probably the most significant K-selection strategy in insects, with females laying only one egg at a time,
albeit in succession, and larvae nursed internally in the mother by special uterine
“milk” glands (Oldroyd 1964). Hippoboscoidea have considerable mouthpart penetrability and styletal dexterity for puncturing the often-hardened integument of
most vertebrate hosts. Perhaps because of thickened and indurated host integument,
the adult mouthparts consist of a short, rigid proboscis with prominent, prestomal
teeth mounted on a labellum used for abrading and cutting through integumental
tissue to feed on upwelling fluids such as lymph and blood (Jobling 1926, 1928,
1929, 1933).
Glossinidae (tsetse flies) are large, hematophagous flies with distinctive mouthparts that are obligately parasitic on vertebrates, including humans (Jobling 1933).
From their blood feeding, glossinids are vectors of the trypanosome Trypanosoma
that cause the debilitating and fatal diseases of rinderpest and nagana in livestock
and sleeping sickness in humans throughout modern, sub-Saharan Africa (Lambrecht
2018). Glossinids consist of a single modern and fossil genus, Glossina. Glossina
contains 15 species that is split into three subgenera, each of which occupies a
savanna, forest or riverine–lacustrine margin habitat. Two Glossina species have
been found in the 34 million-year-old, latest Eocene Florissant Formation in
Colorado, U.S.A. (Grimaldi 1992), and an additional species is known from a 10
million-year-older occurrence at the Enspel locality in Germany (Wedmann 2000).
This biogeographical distribution indicates that Glossina had a widespread distribution in the Northern Hemisphere during the Paleogene, and became restricted to
sub-Saharan Africa in the Neogene. It remains unknown whether Glossina was a
vector for diseases similar to sleeping sickness during its earlier, Paleogene existence (Martins-Neto 2003).
Hippoboscidae (louse flies, keds) consist of 3 subfamilies and 21 genera of obligate parasites of birds and mammals. Hippoboscid fossil history is extremely limited and is comprised of two occurrences. One specimen is from Rott, Rhineland,
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution
Aspöck 2012) that are parasitic on vertebrates, mostly livestock (Cupp et al. 1998;
Salem et al. 2012), and represent probably a single origination of hematophagy
(Schremmer 1961). Feeding in this tribe is accomplished by sharp, prestomal teeth
on the labellum, resulting in surface abrasion of the integument and sponging of
blood and other dermal fluids that pool in the lesions. The Tribe Stomoxyini lacks a
fossil record, but because the clade is a member of the post-Cretaceous Calyptratae
(Grimaldi and Engel 2005a), the lineage likely originated during the Paleogene and
acquired parasitic interactions with large mammals at that time.
The Hippoboscoidea, termed Pupipara in the older literature, consists of the four
constituent lineages of Glossinidae (tsetse flies), Hippoboscidae (louse flies, keds),
Nycteribiidae (bat flies) and Streblidae (bat flies), although the last two lineages are
not monophyletic (Griffiths 1972; Kutty et al. 2001). Hippoboscoid flies are united
by several, common and distinctive reproductive features. The mature female lays a
single mature larva, the puparium, after which the larva immediately pupariates,
emerging from the hardened, last-instar larval skin that encloses the pupa (Ferrar
1987). The emergent, hematophagous adult then locates and parasitizes a suitable
bird or mammalian host. This unique larval development is probably the most significant K-selection strategy in insects, with females laying only one egg at a time,
albeit in succession, and larvae nursed internally in the mother by special uterine
“milk” glands (Oldroyd 1964). Hippoboscoidea have considerable mouthpart penetrability and styletal dexterity for puncturing the often-hardened integument of
most vertebrate hosts. Perhaps because of thickened and indurated host integument,
the adult mouthparts consist of a short, rigid proboscis with prominent, prestomal
teeth mounted on a labellum used for abrading and cutting through integumental
tissue to feed on upwelling fluids such as lymph and blood (Jobling 1926, 1928,
1929, 1933).
Glossinidae (tsetse flies) are large, hematophagous flies with distinctive mouthparts that are obligately parasitic on vertebrates, including humans (Jobling 1933).
From their blood feeding, glossinids are vectors of the trypanosome Trypanosoma
that cause the debilitating and fatal diseases of rinderpest and nagana in livestock
and sleeping sickness in humans throughout modern, sub-Saharan Africa (Lambrecht
2018). Glossinids consist of a single modern and fossil genus, Glossina. Glossina
contains 15 species that is split into three subgenera, each of which occupies a
savanna, forest or riverine–lacustrine margin habitat. Two Glossina species have
been found in the 34 million-year-old, latest Eocene Florissant Formation in
Colorado, U.S.A. (Grimaldi 1992), and an additional species is known from a 10
million-year-older occurrence at the Enspel locality in Germany (Wedmann 2000).
This biogeographical distribution indicates that Glossina had a widespread distribution in the Northern Hemisphere during the Paleogene, and became restricted to
sub-Saharan Africa in the Neogene. It remains unknown whether Glossina was a
vector for diseases similar to sleeping sickness during its earlier, Paleogene existence (Martins-Neto 2003).
Hippoboscidae (louse flies, keds) consist of 3 subfamilies and 21 genera of obligate parasites of birds and mammals. Hippoboscid fossil history is extremely limited and is comprised of two occurrences. One specimen is from Rott, Rhineland,
11 The History of Insect Parasitism and the Mid-Mesozoic Parasitoid Revolution